Neuron Structure and Function

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1 C H A P T E R 4 Neuron Structure and Function PowerPoint Lecture Slides prepared by Stephen Gehnrich, Salisbury University

2 Neurons Vary in structure and properties Use same basic mechanisms to send signals Figure 4.1

3 Neural Zones Four functional zones Signal reception Dendrites and the cell body (soma) Incoming signal received and converted to change in membrane potential Signal integration Axon hillock Strong signal is converted to an action potential (AP)

4 Neural Zones Signal conduction Axon (some wrapped in myelin sheath) AP travels down axon Signal transmission Axon terminals Release of neurotransmitter

5 Neural Zones Trigger zone Figure 4.2

6 Electrical Signals

7 Electrical Signals in Neurons Neurons have a resting membrane potential (like all cells) Membrane potential is negative at rest Neurons are excitable Can rapidly change their membrane potential Depolarization membrane potential becomes less negative Repolarization membrane potential returns to resting value Hyperpolarization membrane potential becomes more negative than resting value

8 Electrical Signals in Neurons Changes in membrane potential act as electrical signals

9 Changes in Membrane Potential Figure 4.3

10 Membrane Potential Factors contributing to membrane potential Distribution of ions across the membrane Relative permeability of the ions Charges of the ions Goldman equation for the calculation of membrane potential (E m ) Em RT F ln P P K K [ K [ K ] ] o i P P Na Na [ Na [ Na ] ] o i P P Cl Cl [ Cl [ Cl ] ] i o

11 The Goldman Equation Em RT F ln P P K K [ K [ K ] ] o i P P Na Na [ Na [ Na ] ] o i P P Cl Cl [ Cl [ Cl ] ] i o E m = membrane potential R = gas constant T = temperature (Kelvin) F = Faraday s constant P x = relative permeability of ion [X] = ion concentration outside or inside membrane

12 The Goldman Equation Em RT F ln P P K K [ K [ K ] ] o i P P Na Na [ Na [ Na ] ] o i P P Cl Cl [ Cl [ Cl ] ] i o Other ions (Ca ++, Mg ++, etc.) are ignored in this simplified form of the equation because their permeabilities are very low

13 Gated Ion Channels Neurons depolarize or hyperpolarize by selectively altering permeability Gated ion channels open or close in response to a stimulus Example: neurotransmitter

14 Gated Ion Channels Channels only allow specific ions to pass through the membrane Ion moves down its electrochemical gradient Only relatively small numbers of ions move across As permeability to a specific ion increases, membrane potential will approach that ion s equilibrium potential (Nernst equation)

15 Changes in Membrane Potential Figure 4.4

16 Neural Zones Signal reception Signal integration Signal conduction Signal transmission

17 Signal reception

18 Signals in the Dendrites and Cell Body Incoming signal Example: neurotransmitter Membrane-bound receptors bind to neurotransmitter Receptors transduce the chemical signal to an electrical signal by changing ion permeability of membrane Change in ion permeability causes change in membrane potential (graded potential)

19 Graded Potentials Vary in magnitude depending on strength of stimulus More neurotransmitter more ion channels open larger magnitude of graded potential Depolarization Na + or Ca 2+ channels open Hyperpolarize K + and Cl channels open

20 Stimulus Strength and Graded Potentials Figure 4.5

21 Graded Potentials Travel Short Distances Conduction with decrement Magnitude of graded potential decreases with increasing distance from opened ion channel Decrement due to: Leakage of charged ions across membrane Electrical resistance of cytoplasm Electrical properties of membrane Electrotonic current spread Positive charge spreads through cytoplasm causing depolarization of adjacent membrane

22 Conduction with Decrement Figure 4.6

23 Signal integration

24 Action Potentials Travel Long Distances Characteristics of Action Potentials: Triggered by net graded potential at axon hillock (trigger zone) Travel long distances along membrane All-or-none Must reach threshold potential to fire Depolarizations below threshold will not initiate an action potential

25 Action Potentials Figure 4.7

26 Integration of Graded Signals Spatial summation Graded potentials from different sites influence the net change Many graded potentials can be generated simultaneously Many receptor sites Many types of receptors Temporal summation Graded potentials that occur at slightly different times influence net change

27 Spatial Summation Figure 4.8

28 Temporal Summation Figure 4.9

29 Graded Potentials vs. Action Potentials Table 4.1

30 Signal conduction Signal in the Axon

31 Action Potentials (AP) Occur only when membrane potential at axon hillock reaches threshold Three phases: Depolarization Repolarization Hyperpolarization Absolute refractory period (ARP) Cell incapable of generating a new AP Relative refractory period (RRP) More difficult to generate new AP

32 Action Potentials (AP) ARP RRP Figure 4.10a

33 Voltage-Gated Channels Change shape due to changes in membrane potential Closed at resting potential Positive feedback Influx of Na + local depolarization more Na + channels open more depolarization

34 Voltage-Gated Channels Na + channels open first (depolarization) K + channels open more slowly (repolarization) Na + channels close K + channels close slowly relative refractory period caused by open K + channels Figure 4.10b

35 Action Potentials (AP) Figure 4.10

36 Na + Channels Have Two Gates Activation gate Voltage dependent Opens when membrane reaches threshold Inactivation gate Time-dependent Closes after brief time

37 Na + Voltage-Gated Channels Figure 4.11

38 Ion Movement Relatively small number of ions move into and out of cell Single action potential has no measurable affect on ion concentrations inside and outside cell Na + /K + ATPase restores concentration gradients following repeated action potentials

39 Voltage-Gated Channels and the AP Figure 4.12

40 Action Potentials Travel Long Distances All-or-none Occurs or does not occur All APs are same magnitude Self propagating An AP triggers the next AP in adjacent areas of membrane without degradation Electronic current spread Charge spreads along membrane Regenerative cycle Ion entry electronic current spread triggering of AP

41 Action Potentials Travel Long Distances Figure 4.13

42 Myelination Vertebrate neurons are myelinated Myelin Insulating layer of lipid-rich Schwann cells wrapped around axon Reduce leakage of charge across membrane Schwaan cells are a type of Glial cell Cells other than neurons that support neuron function

43 Myelination Nodes of Ranvier Areas of exposed axonal membrane between Schwann cells Internodes The myelinated region Saltatory conduction APs leap from node to node APs occur at nodes of Ranvier, and electrotonic current spread through internodes This type of conduction is very rapid

44 Myelination Figure 4.14

45 Unidirectional Signals Action potentials start at the axon hillock and travel towards the axon terminal Up-stream Na + channels (just behind the region of depolarization) are in the absolute refractory period The absolute refractory period prevents backward (retrograde) transmission and summation of APs Relatively refractory period also contributes by requiring a very strong stimulus to cause another AP

46 Information Transfer by AP AP frequency carries information AP frequency increases with stronger stimuli Magnitude of each AP does not change Maximum frequency is limited by the absolute refractory period Mammalian nerves can conduct action potentials per second

47 Action Potential Frequency Figure 4.15

48 Signal transmission Signal Across the Synapse

49 The Synapse Signal transmission from neuron to another cell Synapse Presynaptic cell, synaptic cleft, and postsynaptic cell Synaptic cleft Space between the presynaptic and postsynaptic cell Postsynaptic cell May be a neuron, muscle cell, or endocrine cell Neuromuscular junction Synapse between a motor neuron and a skeletal muscle cell

50 The Synapse

51 Signal Transmission at a Chemical Synapse Figure 4.16

52

53 Amount of Neurotransmitter Released [Ca 2+ ] i is affected by AP frequency More open voltage-gated Ca 2+ channels [Ca 2+ ] i Factors that lower intracellular [Ca 2+ ] i Binding with intracellular buffers [Ca 2+ ] i Ca 2+ ATPases [Ca 2+ ] i High AP frequency Ca 2+ influx is greater than removal [Ca 2+ ] i many synaptic vesicles release their contents high [neurotransmitter] in synapse

54 Acetylcholine Figure 4.17

55 Postsynaptic Cells Postsynaptic cells have specific receptors for neurotransmitters Example: nicotinic ACh receptors Similar to specific hormone receptors on target cells Binding of neurotransmitter to receptor alters ion permeability of postsynaptic cell Change in membrane potential of postsynaptic cell

56 Transmission of Signal Strength at Synapse Response of postsynaptic cell influenced by amount of neurotransmitter in synapse and number of receptors Amount of neurotransmitter Rate of release rate of removal Release determined by frequency of APs Removal determined by Passive diffusion out of synapse Degradation by synaptic enzymes Uptake by surrounding cells Number of receptors Density of receptors on postsynaptic cell

57 Diversity of Neurons

58 Diversity of Neurons All neurons have three functions: Receive and integrate incoming signals Conduct the signal along the neuron Transmit the signal to other cells Neurons differ in their ability to receive incoming signals Different receptors Neurons differ in mechanism of signal conduction and synaptic transmission

59 Structural Diversity of Neurons Figure 4.18a

60 Functional Classes of Neurons Afferent neurons (sensory) Conduct action potentials towards the central nervous system Efferent neurons (motor) Conduct action potentials from the central nervous system to the organs Interneurons Conduct action potentials between neurons in the central nervous system

61 Neuron Classification Based on Function Figure 4.18b

62 Structural Classes of Neurons Multipolar Many dendrites One axon Bipolar One dendrite (may have branches) One axon Unipolar Single process extending from cell body May split to form afferent and efferent branches

63 Neuron Classification Based on Structure Figure 4.18c

64 Glial Cells More abundant than neurons 90% of cells in human brain are glial cells Do not generate or conduct APs Do not form synapses with neurons

65 Types of Glial Cells Five main types of glial cells in vertebrates Schwann cell Forms myelin on motor and sensory neurons of PNS Oligodendrocyte Forms myelin on neurons in CNS Astrocyte Transport nutrients, remove debris in CNS Microglia Remove debris and dead cells from CNS Ependymal cells Line fluid-filled cavities of CNS

66 Glial Cells Figure 4.19

67 Diversity of Signal Conduction Diverse mechanisms of signal conduction Electrotonic Action potentials Saltatory conduction Chemical and electrical synapses Additional diversity in AP physiology: Shape and speed of action potential due to properties of Na + and K + channels Function of channels Number of channels

68 Ion Channel Isoforms Channel isoforms encoded by different genes Voltage-gated K + channels are highly diverse 18 genes encode for 50 isoforms in mammals Voltage-gated Na + channels are less diverse 11 isoforms in mammals Each isoform has distinct functional characteristics

69 Ion Channel Isoforms Table 4.2

70 Channel Density Density of voltage-gated Na + channels affects signal conduction Increased density of channels lowers threshold Increased density of channels shortens relative refractory period

71 Voltage-Gated Ca 2+ Channels Presence of voltage-gated Ca 2+ channels affects AP Open at the same time or instead of voltage-gated Na + channels Ca 2+ enters the cell causing depolarization Ca 2+ influx is slower and more sustained than Na + influx Slower maximal frequency of APs due to longer refractory period Voltage-gated Ca 2+ channels play key role in function of cardiac muscle

72 Conduction Speed of Axons Two ways to increase speed: Myelination Increasing diameter of axon Table 4.3

73 Speed of Among axons Length constant Capacitance Time constant Table 4.3

74 Cable Properties of Axons Similar physical principals govern current flow through axons and undersea telephone cables Current (I) Amount of charge moving past a point at a given time A function of the voltage (V) drop across circuit and the resistance (R) of circuit

75 Cable Properties of Axons Voltage (V) Difference in electrical potential Resistance (R) Rorce opposing flow of electrical current Ohm s law: V = I R

76 Cable Properties of Axons An axon behaves like an electrical circuit Ions moving through voltage-gated channels cause current across membrane Current spreads electrotonically along axon

77 Cable Properties of Axons Each area of axon consists of an electrical circuit Three resistors: Extracellular fluid (R e ) Membrane (R m ) Intracellular fluid (R i )

78 Current Flow In Axons Figure 4.20

79 Voltage Decreases with Distance Change in membrane potential (voltage) during AP decreases over distance due to resistance Conduction with decrement Higher resistance of intracellular and extracellular fluids causes greater decrease in voltage along axon Lower resistance of membrane causes greater decrease in voltage along axon K + leak channels (always open) Some + charge leaks out Number of K + leak channels will affect current loss and voltage decrease along axon

80 Length Constant ( ) of Axons Distance over which membrane potential will decrease to 37% of its original value Variables affecting length constant: Resistance of cell membrane (r m ) Resistance of intracellular fluid (r i ) Resistance of extracellular fluid (r o ) r o is usually low and constant; and is often ignored is largest when r m is high and r i is low r m /( ri ro) r m / r i

81 Length Constant ( ) of Axons Figure 4.21

82 and the Speed of Conduction Electronic current flow decreases over distance Higher allows more electrotonic current flow and faster speed of conduction

83 Axon Membrane Capacitance Capacitance Quantity of charge needed to create a potential difference between two surfaces of a capacitor Depends on the feature of the cell membrane Lipid bilayer Larger area increases capacitance Thickness of insulating layer Greater thickness decreases capacitance

84 Axon Membrane Capacitance Figure 4.20b and Figure 4.22

85 Time Constant (t) Time over which membrane potential will decay to 37% of its maximal value Variables affecting time constant: Resistance of cell membrane (r m ) Capacitance of the cell membrane (c m ) t = r m c m Low r m or c m result in low t Capacitor becomes full faster Faster depolarization Faster conduction

86 Time Constant (t) Figure 4.23

87 Giant Axons Easily visible to naked eye (up to 1 mm diameter) Not present in mammals

88 Giant Axons Figure 4.24

89 Giant Axons

90 Giant Axons Have High Conduction Speed r m inversely proportional to surface area Large diameter axons have greater surface area and more leak channels; therefore low resistance r i inversely proportional to volume Large diameter axons have greater volume; therefore low resistance As axon diameter increases, r m and r i both decrease r m / ri Resistance of cell membrane (r m ) Resistance of intracellular fluid (r i )

91 Axon Diameter and the Length Constant Figure 4.25

92 Myelinated Neurons in Vertebrates Disadvantage of large axons Take up a lot of space which Limits number of neurons that can be packed into nervous system Large volume of cytoplasm makes them expensive to produce and maintain Myelin enables rapid signal conduction in compact space

93 Myelin Increases Conduction Speed Increased membrane resistance Insulators decrease current loss through leak channels, increasing the length constant Decreased membrane capacitance Increased thickness of insulating layer reduces capacitance, decreasing the time constant High length constant and low time constant increase conduction speed Nodes of Ranvier are needed to boost depolarization

94 Diversity of Synaptic Transmission

95 Synaptic Transmission Transfer of electrical signal from presynaptic cell to postsynaptic cell Electrical synapse Gap junction Chemical synapse Chemical messenger crosses synaptic cleft

96 Electrical and Chemical Synapses Figure 4.26

97 Electrical and Chemical Synapses Electrical synapse Chemical synapse Rare in complex animals Common in complex animals Common in simple animals Rare in simple animals Fast Slow Bi-directional Unidirectional Postsynaptic signal is similar to presynaptic Excitatory Postsynaptic signal can be different Excitatory or inhibitory

98 Structural Diversity of Chemical Synapses Figure 4.27

99 Dendritic spines

100 Structural Diversity of Chemical Synapses Figure 4.27

101 Neurotransmitters Characteristics of neurotransmitters Synthesized in neurons Released at presynaptic cell following depolarization Bind to a postsynaptic receptor and cause an effect

102 Neurotransmitters More than 50 known substances Categories Amino acids Neuropeptides Biogenic amines Acetylcholine Miscellaneous (gases, purines, etc.) A single neuron can produce and release more than one neurotransmitter

103 Neurotransmitters (1) Table 4.4

104 Neurotransmitters (2) Table 4.4

105 Neurotransmitter Action Excitatory neurotransmitters Cause depolarization of membrane Excitatory postsynaptic potential (EPSP) Make postsynaptic cell more likely to generate an AP Inhibitory neurotransmitters Cause hyperpolarization of membrane Inhibitory postsynaptic potential (IPSP) Make postsynaptic cell less likely to generate an AP

106 Neurotransmitter Receptor Function Ionotropic receptors Ligand-gated ion channels Fast Example: nicotinic Ach receptor Figure 4.28a

107 Neurotransmitter Receptor Function Metabotropic receptors Receptor changes shape Formation of second messenger Alters opening of ion channel Slow May lead to long-term changes via other cellular functions Figure 4.28b

108 Receptors for Acetylcholine (Ach) Cholinergic receptors Nicotinic receptor Ionotropic Muscarinic receptor Metabotropic Linked to ion channel function via G-protein

109 Receptors for Acetylcholine Figure 4.29

110 Receptors for Acetylcholine Table 4.5

111 Catecholamine (Norepinephrine or Epinephrine ) Figure 4.30

112 Receptors for Norepinephrine Adrenergic receptors Alpha ( ) Several isoforms Metabotropic Linked to ion channel function via G-protein Beta ( ) Several isoforms Metabotropic Linked to ion channel function via G-protein

113 Receptors for Norepinephrine Figure 4.31

114 Adrenergic Receptors Table 4.6

115 Synaptic Plasticity Change in synaptic function in response to patterns of use Synaptic facilitation Repeated APs result in increased Ca 2+ in terminal Increased neurotransmitter release Synaptic depression Repeated APs deplete neurotransmitter in terminal Decreased neurotransmitter release

116 Synaptic Plasticity Post-tetanic potentiation (PTP) After train of high frequency APs there is increased neurotransmitter release Exact mechanism unknown, but believed to involve changes in Ca 2+ in terminal

117 Post-tetanic Potentiation (PTP) Figure 4.32

118 Evolution of Neurons Only metazoans have neurons Other organisms have electrical signaling Algae have giant cells that can generate APs using Ca 2+ activated Cl channels Plants have APs involving Ca 2+ that travel through the xylem and phloem Paramecium can change direction as a result of APs produced by Ca 2+ channels Only metazoans have voltage-gated Na + channels

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